 Bubalus
quarlesi
Mountain anoa |
Taxonomy | Description
| Reproduction | Ecology
| Behavior | Distribution
| Conservation | Remarks
| Literature |
| Taxonomy
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Bubalus quarlesi [Ouwens, 1910].
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Citation: Bull. Dépt. Agric. Indes Néerl., 38:7.
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Type locality: Indonesia, Sulawesi, mountains of C Toradja Dist.
The taxonomic record (above) is taken from Wilson and Reeder (1993). The
taxonomy of the anoas - including how many species there are, and whether
the different species are simply morphotypes - is still uncertain, and as
a result, B. quarlesi is included in B. depressicornis by some
authors (see Whitten, 1987). The mountain anoa belongs to the subgenus
Anoa [Hamilton-Smith, 1827], which some authors argue should be upgraded
to a full generic rank (see Nowak, 1991). B. quarlesi is monotypic,
and has no synonyms (Wilson and Reeder, 1993).
General Characteristics
Similar in form to the lowland anoa (B. depressicornis), the mountain
anoa is the smallest of the extant wild cattle (Buchholtz, 1990). The
body is round and barrel-shaped, and the legs are slender (Buchholtz, 1990).
Body length is approximately 150 cm and shoulder height is typically
70 cm (Groves, 1969; Buchholtz, 1990). The discrepancy in reported
adult weights is marked; nearly a three-fold difference between the two reported
values, and unresolvable due to a lack of published data.
Reported measurements for mountain anoa (Bubalus quarlesi) |
| Source |
Adult Weight |
Head & Body Length |
Shoulder Height
|
Tail Length |
| Buchholtz, 1990 |
150 kg |
150 cm |
70 cm |
24 cm |
| Feer, 1994 |
- |
- |
70 cm |
- |
| Groves, 1969 |
56 kg
(n=1) |
152.4 cm
(n=1) |
69 cm
(n=1) |
- |
The coat of adults, especially females, is often thick and woolly into adulthood
- rather than being grown specially, this appears to be merely a retention
of a juvenile characteristic (Groves, 1969). This woolly coat may be
shed with increasing maturity, but if it is lost, the hair is never as sparse
as in B. depressicornis (Groves, 1969). The color of adults
varies from dark brown to black, with males approaching black and generally
being darker than females (Groves, 1969; Feer, 1994). Compared to B.
depressicornis, B. quarlesi has small and inconspicuous markings
(Groves, 1969). The slender legs are brownish black along their entire
length, and the small white spots which may be present just above the hooves
are often difficult to see (Groves, 1969; Buchholtz, 1990). The groin
region is lighter in color than the back, but is not white (Groves, 1969).
The tail of the mountain anoa is shorter than that of B.
depressicornis, being 14.6-17.8 % of the total length (Groves, 1969).
There are usually no markings on the face, and there is never any crescent
of white on the throat (Groves, 1969). The ears are small and relatively
narrow (Buchholtz, 1990). Both male and female mountain anoa possess
a pair of horns (Groves, 1969; Buchholtz, 1990). The horns are relatively
short and conical, and can be distinguished from those of B.
depressicornis by being round, rather than triangular, in section (Groves,
1969). The horns are smooth, without any marked ridges or external
keel (Groves, 1969; Buchholtz, 1990). Horn length for both sexes varies
between 14.6-19.9 cm (Groves, 1969).
Ontogeny and Reproduction
As with many ungulates, males test the receptiveness of females by checking
hormones present in urine, performing "flehmen" (lip-curl) to expose the
urine to the Jacobson's organ along the roof of the mouth. The
flehmen position is usually held for seven seconds. Males occasionally
flehmen after licking their penis, while females may perform the behavior
to a diverse array of olfactory stimuli. A male will follow a receptive
female, with his head stretched forward at the height of the shoulder and
the neck horizontal - a position which allows the male to lick the female's
vulva to induce urination. Stamping of the forefeet and raising the
neck so that the throat comes into contact with the rump of the female
escalate the pursuit. After a continued sexual parade interspersed
with testing the female's urine, the male places his chin on the rump of
the female as a final test, and then mounts. During copulation, the
head and neck of the male are pressed against the back and shoulders of the
female, with the front legs pressed against her flanks. Throughout
courting, both males and females make shorts moos when excited (Feer, 1994).
The gestation period is reported to be 276-315 days by the National Research
Council (1983), although the species of anoa is not specified. A captive
female at the Krefeld Zoo in Germany had two calves born 13 months apart
(Feer, 1994). Infant mountain anoa are noticeably lighter in color
than adults, being born a golden brown and growing gradually redder and then
darker brown with age (Groves, 1969; Feer, 1994). The life expectancy
for anoas is 20-25 years (National Research Council, 1983).
Ecology
Mountain anoas inhabit primary rainforest, usually in hilly landscapes, at
elevations up to 2,300 meters (National Research Council, 1983; Sugiharta,
1994). They are typically found in dense forest (rather than in adjacent
open meadows), and prefer habitats with a relatively low density of overstory
trees and high diversity of understory vegetation (Foead, 1992 in IUCN, 2004;
Sugiharta, 1994). Mountain anoas typically live near abundant
water sources in areas with low human activity (Sugiharta, 1994).
Dominant overstory plants in areas frequented by mountain anoa in Lore
Lindu National Park (central Sulawesi) include Castanopsis
acuminatissima, Cryptocarya pacifica, Calophyllum soulatrri,
Lithocarpus celebicus, and Sagraea glabra, while dominant
understory plants include Elastostema lineolatum, Carex
verticillata, Axonopus compressus, Panicum trichoides,
and Elaphantopus scaber (Sugiharta, 1994).
Observations of tracks indicate that mountain anoa travel singly or in pairs,
but never associate in larger groups (Sugiharta, 1994). However, Meijer
(1983, in Whitten et al., 1987) reported a herd of five animals on
Mount Nokilalai running past his expedition. A pair of mountain anoa
is most likely to be comprised of a mother and her offspring, or an adult
male and female (Sugiharta, 1994). The National Research Council (1983)
reports that B. quarlesi is active primarily in the morning, seeking
shelter under shade trees during the afternoon. Sugiharta (1994) found
signs of mountain anoa under fallen trees with a diameter greater than 60
cm, in areas under the roots of live trees, and under large overhanging rocks
along river banks, which he suggests are used for cover. One individual
was observed bedded in Gleichenia linearis at the border between forest
and grassland at 0430 hours in Lore Lindu National Park (Sugiharta, 1994).
Anoas will wallow and bathe in pools of water and mud (National Research
Council, 1983).
B. quarlesi feeds on plants in both forest and open areas, including
those created by avalanches and characterized by secondary plant succession
(Sugiharta, 1994). Sugiharta (1994) noted 27 species of plants which
were fed upon by B. quarlesi in Lore Lindu National Park in central
Sulawesi. These included seven grass and grass-like species (Axonopus
compressus, Oplesmenus corpositus, Panicum trichoides,
Paspalum conjugatum, Carex filicina, Carex speciosa,
and Carex verticillata), three ferns (Diplazium repandum,
Neprolepis radicans, and Sphaerostephanos unitus), seedlings
of three tree species (Castanopsis acuminatissima, Elaeocarpus
moribundus, and Calophyllum soulattri), and several forbs
(Calamus sp., Homalomena sp., Elaphantopus scaber,
Ageratum conyzoides, Blumea balsamifera, Desmodium
scorpiurus, Salvia hispanica, Pandanus furcatus, Rubus
moluccanus, Solanum involucratum, Elastostema lineolatum,
Tetrastigma pedunculare, Deparia patersonii, and Globa
strobilifera). E. lineolatum and the three species of fern
are fed upon most frequently (Sugiharta, 1994). Dung samples analysed
by Green (in Whitten et al., 1987) were composed of 38% woody plants,
9% ferns, 16% broad-leaved plants, 17% grasses and sedges, 5% other monocots,
and 13% moss. Anoa are reported to visit mineral-rich hot springs and
salt licks - the National Research Council (1983) reports they may drink
seawater, which might fulfil their mineral needs in areas without licks or
springs.
Behavior
Feer (1994) conducted one of the only behavioral studies of the mountain
anoa, observing a captive pair and their two offspring (one male and one
female, both immature) at the Krefeld Zoo, Germany. Where possible,
his results are generalized here, but because these behavioral observations
were restricted to four individuals they may not be applicable to the species
as a whole.
Defecation and urination are performed separately in B. quarlesi.
Both males and females adopt the same posture when defecating, with
the back hunched, hind legs crouched, and tail raised. Urination postures
differ between the sexes, with males merely moving their hind legs back slightly,
while females lower their rump by bending and spreading their hind legs,
elevating the tail so it is more or less horizontal.
The adult male in Feer's study performed several behaviors which Feer suggested
might serve to reinforce territorial ownership or the dominance hierarchy
within the group. These behaviors included thrashing bushes and
branches with his horns, digging up soil with the horns, and scraping the
ground with the front feet (typically after smelling the area, and followed
about half the time by urinating over the scraped up soil). The hierarchy
between individuals was reinforced with dominant individuals pushing the
forehead or tapping the horns on the flank or rump of a less dominant animal,
resting the chin or cheek on the rump, or placing one (rarely both) forelegs
on the body. On the approach of a dominant animal or in response to
contact, a submissive animal would adopt a posture with the head stretched
low and tilted forward, with the horns pressed close to the neck. A
more extreme submissive behavior involved the less dominant animal lying
down, pressing the neck and chin to the ground.
The aggressive encounters observed were classified as non-contact and direct.
Indirect encounters typically involved posturing and ritualized displays,
with one animal lowering the head so that the muzzle was near the ground
and forehead nearly vertical. If this posture is returned by the other
animal, the two anoas face each other, but rarely approach. This "standoff"
behaviour was seen in 18% of aggressive encounters, and is likely a threat
used to assert a point but avoid an actual fight. In more aggressive
situations, the head is bent further, with the forehead close to the ground
and the horns pointed towards the adversary. Aggressive charges are
directed straight at the adversary, with the head in a normal position (at
the height of the withers) or raised with the ears pointing behind. This
is accompanied by short, soft grunts or moos. If the opponent does
not flee, the charge ends with a sharp blow. This may be followed by
using the forehead to hit the shoulder or flank, or by using the horns to
strike at the opponent's belly or flank with upward strokes. Among
the four anoa in Feer's study, the adult male and female were more likely
to initiate indirect encounters (64%) than full-contact ones (36%), while
the two younger animals used direct contact with much greater frequency (95%).
Aggressive behaviours initiated by the adult male were directed primarily
towards the young male (47%; the remainder split evenly between the mature
and immature females), while the adult female displayed these same behaviours
primarily to the adult male (88%). The agonistic behaviors initiated
by the young male were directed towards the adult female twice as frequently
as the two other animals. The young female rarely initiated encounters
(3% of all observations). Parallel displays - seen in the majority
of the Bovinae - were not observed by Feer, although they may occur in the
presence of two adult males.
Feer (1994) found solitary play in captive specimens in the form of galloping
around or chasing at objects, including birds. Play behaviour with
conspecifics was elicited by galloping towards them with wild leaps, or with
a shaking of the head with a presentation of the horns. This would
lead to play fighting, consists of head-to-head shoving matches with horns
interlocked and heads bent to the ground (and thus facing backwards).
Occasionally, the anoas observed dropped onto their "knees"
(carpal-metacarpal joints) when sparring. While fighting, the animals
occasionally tried to disengage and butt the flanks of their adversary with
their forehead or horns. In doing so, the pair of individuals would
end up parallel to each other, circling around. Play combat was sometimes
interrupted by face-offs, during with the animals stood face to face with
their heads to the ground, and by rapid chases, during which the 'follower'
ran at their partner with wild movements. Most play behaviour was observed
between the two young individuals. Playful advances by a young male
to an adult male elicited a response only rarely, while the adult female
(his mother) responded more playfully.
When excited, either during play or sexual activity, B. quarlesi will
vocalize with brief moos (Feer, 1994).
Distribution
B. quarlesi is endemic to the Indonesian islands of Sulawesi and Buton
[Butung] in southeast Sulawesi Province, occupying an area approximately
5,000 km² (IUCN, 2004). This species appears to be distributed
across most of its known historical range, and is found in several protected
areas, including Lore Lindu National Park (IUCN, 2004).
Countries: Indonesia (Sulawesi) (IUCN, 2004).
Range Map (Localities redrawn from Groves, 1969)
Conservation Status
The mountain anoa is classified as endangered [C1+2a] by the IUCN (2004),
and is on CITES Appendix I (CITES, 2005). B. quarlesi is fully
protected under Indonesian law, although enforcement is difficult (IUCN,
2004). The total population of B. quarlesi is unlikely to be
more than 3,000–5,000 animals, with the number of mature individuals
likely to be less than 2,500 (IUCN, 2004). There has been a decrease
in the range and abundance of this species since the turn of the century;
and there is no doubt that they have declined over the 1980–2000 period
(approximately 3 generations) (IUCN, 2002). The major threats to the
survival of this species are hunting (mainly for meat), introduced pathogens
and/or parasites, and the loss of suitable habitat to agricultural areas,
with recent reports indicating that hunting is by far the most serious (IUCN,
2004).
Remarks
Anoa is the Celebes name for buffalo. Boubalos (Greek)
a buffalo. The Quarles mountain range runs through central and
southwestern Sulawesi. Although called "mountain anoas" these cattle
are not exclusively montane creatures; some have been seen at sea level (Whitten
et al., 1987).
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French
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Anoa de montagne (Buchholtz, 1990, IUCN, 2004)
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Anoa de Quarle (Buchholtz, 1990, IUCN, 2004)
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German
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Berganoa (Buchholtz, 1990)
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Spanish
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Anoa de montaña (IUCN, 2004)
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Literature
Cited
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Flora and Fauna). 2005. Appendix I, II, and III as adopted by the Conference
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Additional Resources
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* Burton, J. A., S. Hedges, and A. H. Mustari. 2005. The taxonomic
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